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Page 1: nv - IPM · O Q m t C F x mO w O Q W D v r k p Uu yQ O q s y x D r n v Q w C i yQ OQ O vO Hw U L o s t Y W I y x Q u QO t w v m O t sRq rw I J C i yQ D tQ U x r k tQ O O v m a D ...
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cation to the Entscheidungsproblem� Proceedings of the

London Mathematical Society �� �������� �� ����� Cor�

rections� ibid�� �� ����� ��������

http���www�turingarchive�org�viewer��id����title��b

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pers on Undecidable Problems and Computable Functions

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�� David Barker�Plummer�Turing machines� Stanford En�

cyclopedia of Philosophy�

http���plato�stanford�edu�entries� turing�machine�

� George S� Boolos and Richard C� Je�rey� Computabil�

ity and Logic� �rd ed�� Cambridge University Press� �����

�� Robin Gandy� The con�uence of ideas in ���� in Rolf

Herken� ed�� The Universal Turing Machine� A Half�century

Survey� Oxford University Press� ����� pp� �������

�� Piergiorgio Odifreddi� Classical Recursion Theory�

North�Holland� �����

�� Emil Post� Recursive unsolvability of a problem of Thue�

The Journal of Symbolic Logic �� ����� ����� Reprinted

in Davis� op� cit�

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Kinetic trajectories are projected onto xx and yy

variables in �gure �� This �gure shows two popu�

lated states� One corresponds to the loosely gath�

ered coil states while the other is the native state

with a high density�

���DQ� �D CU� xOW xi� � O�OH �xrtH O�OH h�Qo �Q�B u�� QO�xrtH x� ��gure� �xtrm �� pw� �xrtH �Ovm QDu�wQ �Q x�rw� CqtHCQ� �a � � swU �x r tH x � swO �x r tH w CU� xOW pYD t swOcoil� O �OH C�aq�� q�L �CU� xOW �� DQt �two states�h�Qo �Q�B pm uwvm � �OwW�t xO�O swU �xrtH �yDv� QO �statesxrtH OvJ �xawtHt w CU� xOt�QO xJQ�Bm� w sHUvt CQwY x�

�s� Qw��t Qo�O �xvwtv Q� R QO �CU�v sy �� ���DQ���The accuracy of the model structures is given by

TM�score� In case of a perfect match to experimen�

tal structure� TM�score would be ��

O�OH C�aq�� �Q �TM�score� �t�Ok C�aq�� swO �xrtH QOp DN t �a perfect match to experimental structure�

�OwW�t O�yvW�B u� �H x� Q� R �xrtH �CU� xOQmThe accuracy of the model structures is measured

by TM�score� which is equal to � if there is a perfect

match to experimental structure�

w O�OH C�aq�� Twmat ��DQD �tra �yxDWwv pmWt u� QDoQR�p�rO u�� x�CU� umtt C�aq�� uOw� �t�Ok w O�OH �CU� �t�OkQ�m xmv�� �Q� �OW��v K �w w� �Q� CU�vW� �w wt �� xOvU� wv xmO��� O�vm�t �wQW �Q O�OH �xrtH x�oQy O�yO s�Hv� �DUQOx� �Q

�xv �� CU� xOW QmP xr�kt QO �q�k xtrm u�� ��� xm O�UQB� OwN R�

pa�i R� Oa� CUQO �Q pai xm OvQ�O Q�Dv� �U�rov� uwDt u�oOvv�wN �

xJ xm O� wo�t k�w QO xrtH Qy �Ovv��� xrtH x�r�OvUt O�yv ��u�oOvv�wN xrtH l� syi �Q� �OyO�t s�Hv� �Q Q�m xJ �UmQo � �CU�J xOW s�Hv� C�r�ai OyO u�Wv xm OvQ�O �rai x� R��vs�Ot uDt uOv�wN OW�� xDW�O OwHw O�yv w pai u�� O�� R �xrY�iuOv�wN Ov��Qi �k �� �OwW�t �k p�ai� uDi�� �Q� wHDUH ��

�OwW�t QDpmWt sy u� syi xrtH�s� Qw��t �xvwtv �Hv�� QO

The smallest URFs �URFA�L�� a � ��nucleotide

�nt� reading frame overlapping out of phase the

NH��terminal portion of the adenosinetrip hos�

phatase �ATPase� subinit � gene has been iden�

ti�ed as the animal equivalent of the recently dis�

covered yeast H��ATPase subunit � gene�

x� �Ov�xreWtQB �r�N �qwtat ��tvy�Q u�O�DU� w u�� wHWv�O� u�QoWywSBx� �Q Cq�kt �xty xm OwW�t u� R� v�t xOW A�J Cq�kt CQFm xwqaC�aq�� u� QDtyt � QU �r�N xm OvQ�wO�t� �yv� �Ovvm xar��t pt�m CQwYOW��v �r�H u�W��Q� xr�kt u�wva Qo � �qwtat �Ovvm �O�B xr�kt l� QO �Q�yxDi�� �� xHwD p��k �yQw�wv w�L xO�mJ Qo � w Ovv�wN�tv �Q xO�mJ�Q xr�kt xm OvQ�o� s�tYD Qo � �DL �Ovvm�tv xar��t �Q xr�kt OW��v ��PHOvvm�t xar��t �Q �WN� �yvD w OvQPo�t �yh�Qo �Q�B�re� R� Ovvm xar��txm CN�UVwN �xr�kt l� uDWwv u��Q��v� �CU� �r�H u�W��Q� QDW�� xmx� �CU� syt �r�N OyO� u�oOvv�wN x� �Q C�aq�� � QU uDi�� u�mt�O�vDU� u� x� w OQ�o�t Q�Qk xO�iDU� OQwt xOQDUo Qw� x� �xr�kt xwqal� Kw w O�rm �OQm lQO �Q u� C�ty� u�wD� O�� R VqD uwO� xm OwW�tQO �Q u�WQv OQwt C�aq�� uDi�� u�mt� �yxOvv�wN x� xm CU� u�� xr�ktuwO� �Q �tW �xr�kt u�oOvv�wN O�y�wN�t Qo � �s�yO� OvQ�O Q�Dv� xm ���H

�O�vm xOQw�Q� �Q �yv� C�Q�Dv� O��� Ovtyi� O�� R VqD

u�oOvv�wN C�Q��Dv�

xrtH l� R u�oOvvwN CQ��Dv

u� �� xm Ovv��� �Q �D�aq�� xrtH �OD�� QO OvQ�O Q�Dv� u�oOvv�wN ��

CU� �xrtH xOvv�wN �Q� xrtH �wv u� QDu�U� �OvQ�O ���vW�Cq�kt QO Qt� u�� �t� Ov�O�t xOvv�wN xm OW�� R�J �xQ�� QO xmA�J u�mt� O�OH �yOQw�DUO �yvD uwJ CU�v Q�PBu�mt� �traxm CU� �D�Lq�Y� w�L �qwtat �tra �xr�kt xwqa x� �OvQ�O�tyiu�U� ��Lr R� xrtH �wv u�twO �OvQ�O �oR�D u�� wHWv�O �Q��t�Ok C�aq�� u�� Q�wty w u�wQ ���DQ� u� QO xm CU� �xrtH�xOW QmP �q�k ��� �vW� ��r�t �� �va� OW�� xDW�O OwHw O�OH wxDWwv�wN �xr�kt l� QO �O��� xtD�N O�OH C�aq�� �� w xOW R�e�uw v�k �O vwW R�e� ��t �Ok� C�aq�� � � O �� � CqtH �xty xOWu�� ��� ��UQB� COwN R� xm CU� u�� xrtH �OD�� uDWwv ��q�pmWt �yv� uOv�wN xm �Dq�kt QFm � QO �s�xOQm �iQat �q�k �Q swyitQO �OvwQ�t Q�m x� �U�vt �xtOkt uwO� �re� O�OH s�y�it CU�

�O�v���t �Q CUO u�� R� �xvwtv �Hv��Samples for ��dimensional projection of kinetic tra�

jectories are shown in �gure �� The coil states are

loosely gathered while the native states can form

a black cluster with extreme high density in ��

dimensional projection plane�

�OwW�t T�UL� �rqDN� swO �xrtH x � pw� �xrtH R� p�kD v� QOp�tDL� x� Q� R h�Qo �Q�B �CU� xOt� �Hm R� �The coil states�

�CU� QDsyi p��k u�oOvv�wN �Q� O�� R

�� �B��B xQ�tW ��

Administrator
Line
Page 17: nv - IPM · O Q m t C F x mO w O Q W D v r k p Uu yQ O q s y x D r n v Q w C i yQ OQ O vO Hw U L o s t Y W I y x Q u QO t w v m O t sRq rw I J C i yQ D tQ U x r k tQ O O v m a D ...

hQo Q�B l� R u�oOvvwN CQ��Dv

O��� h�Qo �Q�B �O� wo� xr�kt �w wt �xQ�� QO �Q xDmv l� �yvD O��� h�Qo �Q�B Qy�� Ov�tH l� �� w OwW �wQW h�Qo �Q�B �rm uwt�t R� �m �L �xrtH ��O��� h�Qo �Q�Bl� C�rtH �O��� xtD�N Oa� h�Qo �Q�B x� p�kDv� �Q� �xrtHR� �yv� �rm OvwQ w OvW�� ��DQt sy �� �yDv� �D �OD�� R� �k�vt CQwY x�h�Qo �Q�Bl� QO xDmv u�OvJ QmP �OW�� O�OH C�aq�� x� �t�Ok C�aq��Oy�wN�t h�Qo �Q�B xm �Q R�J lQO w Ovm p��vO O��� xOvv�wN �Q xJv� syi

�Ovm�t pmWt O� wo�

�CU� xOW x��Q� h�Qo �Q�B l� �Hv�� QO

The enthalpy of hydrogen bond formation between

the nucleoside bases ��deoxyguanosine �dG� and ��

deoxycytidine �dC� has been determined by direct mea�

surement� dG and dC were derivatized at the � and � hy�

droxyls with triisopropylsilyl grpups to obtain solubility

of the nucleosides in non�aqueous solvents and to pre�

vent the ribose hydroxyls from forming hydrogen bonds�

From isoperibolic titration measurements� the enthalpy

of dC� dG base pair formation is����� ��� kcal�mol�

QO �aU xOvU� wv xm �Q �rY� �xDmv u�wD�t pmWt �r�N h�Qo �Q�B u�� R�x� uDt �yDv� w �OD�� QO �enthalpy� �xtrm x� xHwD �� �O�tyi OQ�O u� u���QDy� VwQ Q� R QO �CU� �w wt u�� Q� h�Qo �Q�B RmQtD xm OUQ�t Qv

�O�v���t h�Qo �Q�B �rY� �w wt u�wva x� �Br�Dv� h�YwD �Q�

We have directly measured the enthalpy of hy�

drogen bond formation between the nucleoside bases

��deoxyguanosine �dG� and ��deoxycytidine �dG��

dG and dC were derivatized at the � and � hydroxyls

with triisopropylsilyl groups� these groups serve both to

solubilize the nucleosides in non�aqueous solvents and

to prevent the ribose hydroxyls from forming hydro�

gen bonds� The enthalpy of dC�dG base pair formation

is ����� ��� kcal�mol according to isoperibolic titration

measurements�

�xrtH Q��eD �CU� xOW h�YwD pw� �xrtH QO h�Qo �Q�B pm �w wt q�LO�OH C�aq�� uOQw� �� �CU� xOW s�Hv� hOy wO �� �rY� uDt �pw��h�Qo �Q�B pm xHwD RmQt x� �yv� p�O�D w xrtH �yDv� QO �dC� w �dG�xrtH wO x� �rY� uDt swO �xrtH �Oa� �xrtH �� QDy� ���DQ� Q�QkQ� ���OW�� xDW�O O�m ��D xr�Ut l� wQ Q� �yvD xrtH Qy �D CU� xOW xDUmWC�aq�� xm O� wo�t �Q h�Qo �Q�B R� �xYqN Kw w x� QN� �xrtH q�L

�s� Qw��t Qo�O �xvwtv �Hv�� QO �CU� xOt� u� �OD�� QO �t�Ok

x�r�OvUt O�yv x� �� Q pai xm �Dr�L QO xrtH u�ty x� p�L�O�vm x�ov CU� xOW QDl�ORv

The smallest of the URFs is URFA�L� a � ��

nucleotic �nt� reading frame overlapping out of

phase the NH��terminal portion of the adenosien�

triphosphatase �ATOase� subinit � Gene� it has

been identi�ed as the animal equivalent of the re�

cently discovered yeast H��ATPase subunit � gene�

w �vqw� �yO�yv R� xO�iDU� R� �OQ�O QDW�� uR�wD O�OH �xrtHnvy�ty lJwm ��B �� nQR� �QU �O�vm ��vDH� x�Dwm �ypwaitCUQO xm O�W�� xDW�O x�Dwm O�yv l� xm CU� QDy� �r�N �CU�v

�Ov���� �vqw� pwait w pai u� R� Oa�

xDmv l� wQ O�m ��D �Q� �yvD xrtH Qy xm OvQ�O Q�Dv� u�oOvv�wN ��

Q� R �xrtH wO �xU��kt �� �CU� xOW xOQw� xrtH �yDv� QO xm OW���OvQ�O O�m ��D �Dw�iDt �yR�J Q� xm O�v��� O�v�wD�t

URFA�L has been identi�ed as the animal equiv�

alent of the recently discovered yeast H��ATPase

subunit � gene�

The recently discovered yeast H��ATPase subunit

� gene has a corresponding animal equivalent gene�

URFA�L�

xOW hWm ��Q�N� xm CU� �vS �xQ�� QO pw� �xrtH xm CU� K �wO�m ��D C�v�w�L QO u� pO�at uS wQ swO �xrtH xm �r�L QO CU�

�OQ�O

�O�v���t �Q Qo�O xvwtv Q� R QO

The enthalpy of hydrogen bond formation between

the nucleoside bases ��deoxyguanosine �dG� and ��

deoxycytidine �dC� has been determined by direct

measurement�

O�m ��D �direct measurement� wQ xrtH u�� xm OUQ�t Qv x�Twmat �� �OW�� xOw� u�� xOvU� wv hOy xm CU� O�a� �t� �OQ�O

�OwW�t Q�QkQ� u� QO QDy� uR�wD xrtH �RH� ��DQD uOQm

We have directly measured the enthalpy of hydro�

gen bond formation between the nucleoside bases

��deoxyguanosine �dG� and ��deoxycytidine �dC��

QU� pmWt R� u� QO u�vJty �CU� QDxO�U w QDx�Dwm O�OH �xrtH�CU� xOW ��vDH� �lJwm ��B w nQR�

�� ���� Q�y�

Page 18: nv - IPM · O Q m t C F x mO w O Q W D v r k p Uu yQ O q s y x D r n v Q w C i yQ OQ O vO Hw U L o s t Y W I y x Q u QO t w v m O t sRq rw I J C i yQ D tQ U x r k tQ O O v m a D ...

pmW w pwOH R u�oOvvwN Q��Dv

w O�it QOkJ x r�kt l � Ovtyi � xmv �� �Q � pwHa u�oOv v�wN ��a �u ��Q �� v � �O vwQ�t �ypwOH w �ypmW d�QU x � ��t �k DUt CU� � r�H�UQ w �� wo �i�m �xR�Ov� x� Q� w�YD w �ypwOH KQW xm CU� syt �r�N��a� OvJQy� �OW��v uDt uOv�wN x� R��v �yv� uO�tyi �Q� �D OW��u�� QO �OvQ�O Q� w�YD w pw�OH KQW �Q� �Q OwN X�N ���w CqHt R�xm �Hv� R� ��OwW xaH�Qt u�oOvU� wv �Q� xrHt ptar�wDUO x� O��� OQ�wt�t�Ok C�aq�� xm s� Q�O Q�Dv� s�v�wN�t CU�Q x� AJ R� �Q pwOHp�Ft �Q� �OvwW xOvQw� CU�Q CtU QO O�OH C�aq�� w AJ CtU QOCw�iD �yuwDU �H �ki �yv� QO xm Ov�xOW xOQw� Q� R QO � w � pwOH wO

�OQ�O

Table �

Temp��C� Time

��

�� �

�� �

�� ��

�� ��

Table �

Time Temp��C�

��

� ��

� ��

�� ��

�� ��

u�tR � � �t uwJ CU� � pwOH R� Q DxO�U �r�N � pwOH uO v�wNu�� pw�OH �Q� Qo�O uwv�k �s�DUy �vW� pkDUt Q�eDt l� u�wva x�xm �vat u�� x� OvwW xOQw� pwOH �yDv� QO C�aq�� u� QDy� xm CU�u � QN� QO � � CU�Q CtU uwDU QO O �� � G ��D v u � Q D�r�H w u � Q DO�itu�� x� w xO�UQ uOv�wN �Q�m �yDv� x� xOvv�wN �Hv� QO �OvwW x��Q� Q�UOvJ CkO Q� R p�Ft QO �OQ�o�t Q�Qk xOW x��Q� G��Dv Q�F��D CLD ��DQDx��Q� �Q �vwvm �xar��t G��Dv Q�U u� QN� �CU� xOW xU��kt sy �� VwQ

�Ovm�t

Benchmark SALIGN Lindahl PROSPECTOR � LiveBench �

Method Alignment MaxSub MaxSub MaxSub

SPARKS ����� ���� ��� ����

SPARKS� ���� ���� ��� ���

This work � � � ���� �� ����

Cvwi R� xm CU� u�� OwW C��aQ O��� �ypmW QO xm R�J pk�OL�s�qa w �yQwLt O�Oa�� �y�UJQ� �xty �Q� Helvetica x��U w nQR�xO�iDU� uwO � �O � Qw�� � pmW QO �Q x�L� v u � Q Dsyt �ki �OwW xO�iDU�xv� Ry nvQ R� xO�iDU�� O�yO x��Q� Kw w QFm �OL �� �Q �yQ�Owtv nvQ R�

��OQ�O

Large earthquakes along a given fault segment do

not occur at random intervals because it takes time to

accumulate the strain energy for the rupture� The rates

at which tectonic plates move and accumulate strain

at their boundaries are approximately uniform� There�

fore� in �rst approximation� one may expect that large

reptures of the same fault segment will occur at ap�

proximately constant time intervals� If subsequent main

shocks have di�erent amounts of slip across the fault�

then the recurrence time may vary� and the basic idea

of periodic main shocks must be modi�ed�

�xQ� � QO �L �w �� D �U v CQwY x � swO w pw� CqtH p� Ft u �� QOQO pw� �xrtH �t�Ok C�aq�� xJQo OvDUy Q�Wi �oDW��v� �nvy��x� xOvv�wN swU �xrtH x� uO�UQ �� �CU� xOWv xOQw� swO �xrtH �OD��

�Ovm�t so �Q �r�t �DL�Q

�OW�� xOvU� wv QvOQwt �r��t R� QDuWwQ KQW Ov�wD�t Q� R h�Qo �Q�B

Large earthquakes along a given fault segment do

not occur at random intervals because it takes time

to accumulate the strain energy for the rupture� The

rates of strain accumulation at the boundaries of tec�

tonic plates are approximately uniform� Therefore�

nearly constant time intervals �at �rst approximation�

would be expected between large ruptures of the same

fault segment� However� the recurrence time may vary�

the basic idea of periodic main shocks may need to

be modi�ed if subsequent main shocks have di�erent

amounts of slip across the fault�

�CU� xrRrR O�ONQ wQ �vWwQ x� O�OH h�Qo �Q�B �rY� RmQtD q�LKQ�t �q�k xm �Q �t�Ok C�aq�� xOW xO�Wm �N �yv� Q� R xm Qw�Ux��Q� h�Qo �Q�B syi pmWt u� QDoQR� xm CU� K �w �OvyO�t u�Wv xOWuOQm QDu�U� �Q� x�Q u� QDy� �Ow� O�OH w s�Ok C�aq�� ��re �H xOWC�aq�� x� �t�Ok C�aq�� R� Q�DWwv OvwQ xm CU� u�� u�oOvv�wN �xar��t

�OW�� O�OH

xmr� Ovm V��tR� �Q u�oOvv�wN lQO �xwk xm CU�v u�� xr�kt l� hOyO�v�wD�tv �tW �OyO u�Wv Vy�oO�O K� QY u��� QO �Q xOvU� wv ���v�wD O���O��� xmr� �O�vm VvRQU u�Dxr�kt syi QO �v�wD�v p�rO x� �Q u�oOvv�wN

�O�vm VvRQU u�DQv OQwt s��B p�kDv� QO �t�m �v p�rO x� �Q u�DOwN

�� �B��B xQ�tW ��

Page 19: nv - IPM · O Q m t C F x mO w O Q W D v r k p Uu yQ O q s y x D r n v Q w C i yQ OQ O vO Hw U L o s t Y W I y x Q u QO t w v m O t sRq rw I J C i yQ D tQ U x r k tQ O O v m a D ...

xr�kt l� �Q� Ovv�wD�t �DNU x� u�Qw�O �okqa�� p�rO x� xrHt��Ovvm �O�B Qw�O

�O�vm x�HwD w O�yO� K� wD �qt�m �Q u�DQ�m R� xrLQt Qy w QDt�Q�B Qy ��

j�kO K� wD �OvQ�Ov Ckw C���RH �xQ�� QO uOQm Qmi �Q� u�Qw�O�O�vm�t xJ O�v�O�t �tW xmCU� u� �xOvyOu�Wv �yQDt�Q�B w �yOv��Qi

�D CU� xOW x��Q� �i�m C���RH �� R�J xty ��� xm O�UQB� u�DOwN R� ���D���RH I�y R� �xv �� Ovm O�rwD R�� �Q �tW G��Dv Ov�wD� Qo�OXNWQDCL�Q ��xOvv�wN Qy ��� Qw�O l� xm QOkQy �O�vmv Qw�a QUQU�Q �tW �xr�kt xm OQ�O QDW�� p�tDL� Ovm O�rwD R�� �Q �tW Q�m Ov�wD��yv� O��� �tW �t� Ovvm�tv O�rwD R�� �Q �tW Q�m k�w QO u�Qw�O �OQ�PB�u�� O��xO�O x��Q� xm �L� wD T�U�Q� Ovv�wD�t xm O�vm Oa�kDt �Q

�OvyO s�Hv� �Q Q�m

xiYv xv �� pt�m CQwY x� �Q j�kLD �OW�� xOvvmv�k u�DQ�m O�vm �aU ��

V��tR� u�OvJ �� �Q �rY� �w wt u�mt� OL �D �O�yO s�Hv� �� xQ�mxm O�U� wv� p�mw l� �x�a�iO x� x��W �Q OwN �xr�kt �O���tR��� �vt ��

�Ovm C��F� �t�y�� w lW I�y uwO� �Q OwN waO Oy�wN�t

wHDUH xr�kt uDWwv u�tR QO �O�yO ��HQ� syt C�ar��t �xty x� ��

�O�yO s�Hv� ��DQt �yQ�m QO �rt�m

xr�kt Q�DN�U

�yptar�QwDUO �yVwQ xtOkt xO�mJ u�wval� pt�W �tra �xr�kt Qyw QmWD w Q�OkD EL� G��Dv �experimental procedures� V��tR�u�oOvv�wN xm OvwW�t xDiQo Qv QO Qwvt u�� x� �yVN� u�� �CU� H�QtCUQO�vVN� QO�r��t uO�O Q�Qk �Ov���� QD� QU �Q OwN Qv OQwt�r��tuDN�t� QO u�� wHWv�O u�� QO G��Q x��DW� l� �Ovm�t G�o �Q u�oOvv�wN

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�OyO�t K�HQD �Q u� xr�kt u�� �xOvU� wv xm CU� �rmW q�� pmWu�Qo�O C�ar��t G��Dv w O�vm xO�iDU� ODtt �w�N R� u�DOwN Q�m �Q�u�� Cw�iD u� QDW�� uO�O u�Wv �Q� �yO�tv �O�yO u�Wv u�J�N �� �Qx� �Q Y w X �yQwLt u�wva �Ov�xOW �DQt u���B x� q�� R� �y�vLvt

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�OUQ� pk�OL x� Q�m uOw�v pt�m pFt �r�qO x� u� uOW

�s�vm xOQw�Q� �Q �r�tDL� u�Qw�O �xDU�wN Qw�J

�CU� Q�Qk u�� R� u�Qw�O C�� Q �rH ��U�U� pwY�

� � u� oOv v�wN x � O�y�wN�t xJ v� u�w va x � �Q �rY� �w wt l � ��

OwW x��Q� xr�kt l� QO xO�� u�OvJ Qo � �O�vm u��aD O�� wo� u�Qw�O�OvwW�t soQOQU u�oOvv�wN

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Qw�O �xkqa w xHwD O�v�wD�tv Qo � �O�vm ��NDv� u�Qw�O Qv �rH �Q�uqw�Ut �y�o� �O�W��v xr�kt A�J Qmi x� CU� QDy� O�vm �rH �Q

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CU�v syt OvwW hPL O��� G��Dv u�� �OwW�t u�oOvv�wN uOW G�o Ea���O��xOQm VqD �yv� uOQw� CUO x� �Q� QOkJ xm

u�wva

�v�wva xm CU� u� Ckw OW XNWt xr�kt �rY� �w wt xmv� R� TBQ�F��D �� �tW G��Dv �tW �yVwQ Ov�wD�t u�wva �O�vm u��aD xr�kt �Q��xrtH l� QO xr�kt l� R� CU� CQ��a u�wva �Ovm d�r�D �Q �tW G��Dvu�wva p�Ft �Q� �O� Q�Po� u�wva QO �Q C�aq�� u� QDxOvvm �we� w u� QDsyt

�Steric restrictions in protein folding� an alpha�helix

cannot be followed by a contiguous beta�strand�

u�wva �rw �Ovm�t d�r�D �Q G��Dv�Interpreting the folding kinetics of helical proteins�

u�wva �O� wo�t G��Dv Q�F��D �xQ�� QO�Native proteins are surface�molten solids� Applica�

tion of the Lindemann criterion for the solid versus liq�

uid state�

xm O�W�� xDW�O xHwD �OQ�O OwN QO �Q G��Dv Q�F��D w �yVwQ wO Qy�Native proteins are surface�molten solids�

�D OW�� X�N w s�a u�tRty u�wva xm CU� sRq �CU� G��Dv R� Q�UiD�O�vm �O�B QDW�� u�oOvv�wN

xtOkt

�wQW �Q xtOkt O�v�wD�t OW u��aD xr�kt u�wva w �rY� �w wt �Dkw�� ��DQt Cq�kt �xty Qw�tH O�yO s�Hv� O��� xm R�J u�rw� �O�vm�Q ��DQt Cq�kt u� QN� �xty xm O�vm wHDUH QOkv� �CU� �rY� �w wt�xty x� �O� Q�O Q��DN� QO �Q RwQx� C�aq�� �xty xm O� wW u�t�t �O�������HQ� �tW x� sy �yv� O�yOv ��HQ� �yv� x� Qo � �O�yO ��HQ� syt Cq�kt�Q Q�m x� �tW pw� Ovm O�vDU� �tW Q�m x� �Um O�y�wN�t Qo � �O�O Ovy�wNv��a� xmv�� p�tDL� O�vm O�vDU� QDW�� Cq�kt x� QOk Qy �O�yO� ��HQ�OwW�t QDW�� OvyO ��HQ� u� x� w Ovv�wN� �Q �tW �xr�kt �yv� u�oOvU� wv R�xHwD QDW�� CU� xO�O ��HQ� �yv� Q�m x� xm �Dq�kt x� u�YYNDt uwJCkO �� �Q Qo�O Cq�kt uO�O ��HQ� QO x��DW� R� R�yQB CyH �Ovvm�t

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�xtOkt �rm OvwQ uwJ CU� xrtH u�rw� uDWwv �Q� xrtH u� QDCNUu�wva x � �Q xrtH u� rw� xm CU� u �� ut VwQ �Ovm�t u��a D �Q �tWC�Lq�Y� R� ��a� h�Qo �Q�B u�rw� QO ��DQD u�� x� �syO �� Q xr�kth� QaD �yv� u� QD�x��B �� u� QD�twta R� �wQW �� u�wva QO xDiQ Q�m x��OW�� �w wt R� �xv�OkDvt �r�tH� �UQQ� O��� swO h�Qo �Q�B �OwW�t

T�U�Q� �� O��� QDt�Q�B Qy CU� ���yQDt�Q�B pt�W xO�iDU� OQwt VwQ Qo �xOQDUo R�Uxv�y��uwtR� �� w �m� R�i�� �� Q pqODU� �r�k OQ� Q�mxO�O KQW �yQDt�Q�B Q��eD Q�F��D O��� OQ�Ov OwHw x�HwD u�mt� Qo � �OwW x�HwD�yVwQ VN� �OwW x��Q� EL� �� G��Dv VN� QO O��� �ak�w G��Dv� OwWOQ�wt u�� R� l� I�y Qo � ��OW�� �yQDt�Q�B Q�F��D R� �LQW �Q�O O��� �yvD�CU� xOw� xv� RyQB �r�N xmv��� OwW x��Q� u� p�rO O��� OW�� xOWv s�Hv�u� x� Oa� C�ar��t QO �D xOW xDN�Ov� j� waD x� �CU� xOw� Q�u�tR �r�N

��OwW xDN�OQBuOwtR� �Q� �irDNt �yuwtR� O��� u�vJty O�OH VwQ �x��Q� s�ovyu�mt� OL �D xm CU� xOvvm Oa�kDt �DQwY QO O�OH VwQ �O�vm �L�Q� u�p�tDL� O�vm �L�Q� �� xOQm �O�B QDW�� �yuwtR� xJ Qy �OwW xOwtR�

�Ovvm xO�iDU� u� R� u�Qo�O w xOW xDiQ�PB �tW Q�m xm OQ�O QDW����� �� �O�UQB� u�DOwN R� �Q Cq��wU u�� �yVwQ VN� u� wOD R� TBOv�wN� �Q VN� u�� �Um Qo � �� �Ov�xOW h� QaD O�OH C�Lq�Y� �xtyO�� x� �OQ�o�t Q�Qk VQ��DN� QO Q�m pm O�rwD R�� �Q� �i�m C�aq�� ���Qo � sOQt �O�vmv �iNt �Q xOW xO�iDU� Q�v��t �� w sr I�y xm O�W�� xDW�Oxm O�vmv �aU RoQy �OvwW�t CL�Q�v Ovvm O�rwD R�� �Q �tW G��Dv Ovv�wDv�tW R� �yv� R� �r�N xm OvQ�O OwHw O�� R Vwy�� �yu�Uv� �O�vm �rkDO�vm�t Qmi Qo � �OwW�t swrat OwR �� Q�O O�vm �rkD Qo � �OvDUy QDVwy��A�J VRQ� u�DG��Dv Ovm Q�QmD �Q �tW Q�m xm Ovm�tv VqD Tm I�y xm

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�Ovm XNWt hrDNt �yuwtR� �� w hrDNt

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�gurations of helices and sheets� As a result� de�ning

the boundaries between helix� sheet� and coil is prob�

lematical and a signi�cant source of discrepancies be�

tween diferent methods�

Inconsistent assignment of secondary structures

by di�erent methods highlights the need for a crite�

rion� or a benchmark� of �standard� assignments that

can be used to assess and compare assignment meth�

ods� One possibility is to use the secondary structures

assigned by the authors who solved the protein struc�

tures� STRIDE� in fact� has been optimized to achieve

the highest agreement with the authors� annotations�

However� it is not clear what is the criterion used for

manual or automatic assignment of secondary struc�

tures by di�erent authors� Another possibility is to treat

the consensus prediction by several methods as the gold

standard� However� there is no obvious reason why each

method should weigh equally in assigning secondary

structures� or which method should be used in consen�

sus� Other used criteria include helix�capping propen�

sity� the deviation from ideal helical and sheet con�gu�

rations and structural accuracy produced by sequence�

to�structure alignment guided by secondary structure

assignment�

In this paper� we propose to use sequence�alignment

benchmarks for assessing secondary structure assign�

ments� These benchmarks are produced by �D�structure

alignment of structurally homologous proteins� Instead

of assessing the accuracy of secondary�structure assign�

ment directly� which is not yet feasible� we compare the

two assignments of secondary structures in structurally

aligned positions� We assume that the best method

should assign the same secondary�structure element to

the highest fraction of structurally aligned positions�

Certainly� structurally aligned positions do not always

have the same secondary structures� moreover� di�erent

structure�alignment methods do not always produce the

same result� Nevertheless� this criterion provides a mean

to locate a secondary�structure assignment method that

is most consistent with tertiary structure alignment� We

O��� h�Qo �Q�B u�� �CU� xr�Ut l� ��RH pL �� pL �xQ�� QO �rY� �w wtxm �Q ���yVr�J �� CqmWt w xOQm xQ�W� OwHwt �xOWv pL �xr�Ut x��Q O�yvW�B pL x�Q swU h�Qo �Q�B TBU �OyO KQW CU� u� pL R� v�t�tW �OyO�t x��Q� xOt� xr�kt QO xJv� R� QYDNt KQW w xOQm �iQatQ�YDN� x� �Hv�� QO �Q �yv� �r�tDL� C�Q�F��D w xOt� CUO x� G��Dv O�v�wD�t

�CU� xOW xOQw� �xvwtv Q� R QO �O�vm u���

Assessing secondary structure assignments of protein

structures by using pairwise sequence�alignment bench�

marks�

The secondary structure of a protein refers to

the local conformation of its polypeptide backbone�

Knowing the secondary structures of proteins is essen�

tial for their structure classi�cation���� understand�

ing folding dynamics and mechanisms���� and dis�

covering conserved structural� functional motifs ���

Secondary structure information is also useful for se�

quence and multiple sequence alignment��� struc�

ture alignment����� and sequence to structure align�

ment �or threading������� As a result� predicting

secondary structures from protein sequence contin�

ues to be an active �eld of research� ��� �fty six

years after Pauling and Corey��� �rst predicted

that the most common regular patterns of pro�

tein backbones are the ��helix and the ��sheet�

Prediction and application of protein secondary structures

rely on prior assignment of the secondary�structure el�

ements from a given protein structure by human or

computational methods�

Many computational methods have been developed to

automate the assignment of secondary structures� Ex�

amples are DSSP� STRIDE� DEFINE� P�SEA� KAKSI�

P�CURVE� XTLSSTR� SECSTR� SEGNO� and VoTAP�

These methods are based on hydrogen�bond patterns�

geometric features� expert knowledge� or their combi�

nations� However� they often disagree on their assign�

ments� For example� disagreement among DSSP� P�

CURVE� and DEFINE can be as large ad ���� More

beta sheet is assigned by XTLSSTR and more pi�

helix by SECSTR than by DSSP� The discrepancy

among di�erent methods is caused by non�ideal con�

� ���� Q�y�

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potential� Also� an analogous complex phase diagram

is found in simulations of LJ clusters� The present re�

sults for square�well homopolymers may well be found

in more realistic homopolymer models and even in real

polymers�

�OR�OQB�t umtt Q�U�iD Qo�O �UQQ� x� h�Qo �Q�B u��

xO�mJ

KQW pt�W �qwtat xO�mJ �O�U� wv� �Q xO�mJ O��� xr�kt uOW xO�t� R� TBOQwt CU� Q�Qk xm �xr�Ut �u�wva � � �� DQt� �w wt �xv�tR C�ty��CU� G��Dv Q�F��D w �vat C�ty� �tW OQm� wQ �ov�o� OQ�o Q�Qk �UQQ�

�CU� xOW x��Q� QF�wt �xO�mJ l� R� �xvwtv �Hv�� QO

How to make an objective assignment of secondary

based on a protein structure is an unsolved prob�

lem� De�ning the boundaries between helix� sheet�

and coil structures is arbitrary� and commonly ac�

cepted standard assignments do not exist� Here� we

propose a criterion that assesses secondary�structure

assignment based on the similarity of the secondary

structures assigned to structurally aligned residues

in sequence�alignment benchmarks� This criterion is

used to rank six secondary�structure assignment meth�

ods� STRIDE� DSSP� SECSTR� KAKSI� P�SEA� and

SEGNO with three established sequence�alignment

benchmarks �PREFAB� SABmark and SALIGN��

STRIDE and KAKSI achieve comparable success rates

in assigning the same secondary structure elements to

structurally aligned residues in the three denechmarks�

Their success rates are between ���� higher than those

of the other four methods� The consensus of STRIDE�

KAKSI� SECSTR� and p�SEA� called SKSP� improves

over assignments over the best single method in each

benchmark by an additional ��� These results support

the usefulness of the sequence alignment benchmarks as

the benchmarks for secondary structure assignment�

pLx�Q �xQ� � QO swU �x rtH �OvyO�t KQW �Q x r �Ut pw� �x rtH wO�� O��� xO�mJ �CU� xOt� G��Dv CqtH u�� �xt�O� QO �CU� O�yvW�B

�O��� u���B CU� �r�t R� �xYqN w�L xm �xrtH

suggest that this approach provides an objective evalu�

ation of secondary structure assignment methods�

swO �yQ� DN�U � �� � RQ� �Q � �WwQ x r� kt u�w va p� Ft u �� QOswO Q� DN�U R� �i � Qa D � � h�Qo �Q� B u� rw� �Ovm�t d�r� D �Q xOW u��a DC�ty� �xQ� � QO h�Q o �Q� B p m TBU �OwW�t �wQW �u�w va � � �� DQt��yVwQ x� xm CU� �r�kDv� xrtH u� QN� �OvR�t hQL swO �yQ�DN�UVN� u�twO� OwW�t QHvt swO �yQ�DN�U ���� RQ� �Q� �D��U�Lt�yDv� QO �computational methods� xm O�W�� xDW�O xHwD ��u�wvaRmQt D �CU� swO h�Qo �Q� B �OD �� � � �� � DQ� O�H �� w O�m �� D �Q � xrtHCU� � D� �U� L t �yVwQ x m OwHw t �x r �U t wQ Q � swO h�Q o �Q� BG � QO D x � �computational methods� �t �Ok C�aq�� �OW� ��tR� �Q EL� swU h�Qo �Q�B �OwW�t �their disagreement� x� p�O�Dbenchmark for� x� G� QOD x� ��t�Ok C�aq��� �disagreement�TBU �OyO�t Q��eD xrtH u�rw� QO �O�OH C�aq��� �assessment

Q��at sQ�yJ h�Qo �Q�B �OvwW�t �iQat xv�tR u�� QO xOW s�Hv� �yQ�m�OvR�t hQL O�OH Q��at ���Rt �xQ�� QO w xOQm �iQat �Q xOW O�yvW�BKQW �Q G��Dv Q�YDN� x� �CU� xOWv xO�O u�Wv �Hv�� QO xm� sHvB h�Qo �Q�B�Q� �y�HwD �w wt R� �rm �i�YwD ���yvD x� O��� xtOkt Qy �OyO�tO��� u�oOvv�wN xtOkt uOv�wN R� TB �OW�� �yv� Q�F��D w G��Dv O�yvW�B Q�m�OvW�� xDW�O �yv� QF� w G��Dv u�vJty w �tW OQm� wQ w xR�ov� R� �vWwQ QwYD

EL�

�Q EL� uDWwv �y�r�N �O�U� wv� �Q xr�kt VN� u� QN� O��� �tW uwvm �R� G��Dv uOQm �OH QO �re� u�� wHWv�O �Ovv�O�t �yVN� �x�k� R� QDCNUumtt Q�U�iD Qo�O xwqa x� �OvDU�v jiwt Q�oxH�Dv w �yOt��B �yv� Q�U�iD

�OvQ�o�tv Qv QO �Q�OwW�t R�e� �yv� Q�U�iD w xOt� CUO x� G��Dv Q� QwQt �� �wN EL�Q�F��D xOW s�Hv� hrDNt �yuwtR� VN� u�� �Q� �wN EL��t Qo�O�yQ�m xOWv pL p��Ut C�ar��t Qo�O �� xU��kt G��Dv Q� �yQDt�Q�B Q��eDR� EL� VN� R� �xvwtv �Hv�� QO �CU� xOv�� �yQ�m �� s�Hv� p�L QO

�CU� xOW xOQw� xr�kt l�One question about the complex homopolymer phase

diagram presented here is whether it is caused by the

discontinuous feature of the square�well potential� We

connot give a direct answer because the DMD simula�

tion is required to obtain well�converged results for the

thermodynamics� However� the critical phenomena pre�

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Membrane properties and the balance between

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The eect of an information integration task training

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Brain connectivity analysis in fMRI

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rat

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scale device application and second generation

superconducting cables

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measurements

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